Dvulikiaspis Temporal range: Lochkovian, | |
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Interpretive drawing of PIN 1271/2, the holotype of Dvulikiaspis menneri | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Chelicerata |
Clade: | Dekatriata |
Order: | † Chasmataspidida |
Family: | † Diploaspididae |
Genus: | † Dvulikiaspis Marshall et al., 2014 |
Type species | |
†Dvulikiaspis menneri Novojilov, 1959 |
Dvulikiaspis is a genus of chasmataspidid, a group of extinct aquatic arthropods. Fossils of the single and type species, D. menneri, have been discovered in deposits of the Early Devonian period (Lochkovian epoch) in the Krasnoyarsk Krai, Siberia, Russia. The name of the genus is composed by the Russian word двуликий (dvulikij), meaning "two-faced", and the Ancient Greek word ἀσπίς ( aspis ), meaning "shield". The species name honors the discoverer of the holotype of Dvulikiaspis, Vladimir Vasilyevich Menner.
Its prosoma (head) was subquadrate (almost square) to parabolic (nearly U-shaped), with (bean-shaped) to subovate (nearly oval) eyes and surrounded by a marginal rim. The abdomen was composed by a fused buckler and a postabdomen that occupied most of the body length, while the telson (the posteriormost division of the body) was small and semicircular in shape. The appendages (limbs) were uniform. The sixth and last pair of them had a paddle-like shape and was placed in front of the midpoint of the prosoma. The largest specimen was 2.64 centimetres (1.04 inches) long.
The first fossil was described in 1959 as a new species of the eurypterid Stylonurus , while the other two were discovered in 1974. It would not be until 2014 when D. menneri was recognized as a chasmataspidid genus, being placed in the family Diploaspididae. However, Dvulikiaspis was similar to Loganamaraspis and especially Hoplitaspis , with which it could form a new separate family of chasmataspidids.
Like the other chasmataspidids, D. menneri was a small arthropod. The biggest specimen, PIN 5116/1, reached a total length of 2.64 centimetres (1.04 inches). This size is, however, notable among diploaspidids. [1]
The prosoma (head) was subquadrate (almost square) to parabolic (nearly U-shaped), with a carapace (dorsal plate of the prosoma) rather vaulted and somewhat rounded anteriorly and laterally and flat medianly and posteriorly. It was surrounded by a marginal rim that tapered backwards to the sides. The largest preserved prosoma had a length of 0.5 cm (2 in). The eyes were large and reniform (bean-shaped) to subovate (nearly oval), placed between the sides and middle of the carapace. The ocelli (simple eye-like sensory organs) were positioned in the center of the prosoma. [1]
As in the rest of the chasmataspidids, its opisthosoma (abdomen) was composed of a preabdomen (segments 1 to 4) and a postabdomen (segments 5 to 13), in Dvulikiaspis with very slight first order differentiation (that is, both parts little separated from each other). The segments in general were wide rectangles with almost straight boundaries, narrowing slightly posteriorly. The microtergite (the short first tergite, dorsal half of the segment) was conformed to the posterior margin of the prosoma. The second to fourth segments were merged into a weakly expressed "buckler", which was wide rectangular with rounded lateral edges and narrow-angled shoulders (anterolateral "extensions" of the buckler). In the postabdomen, segments 11 to 13 were somewhat longer than the previous ones. Each tergite carried an articular surface (joint) on its front, interpreted to represent arthrodial membranes (plane joints), [1] which were a smooth and flattened facet, with a slight posterior ridge. These joints are identical to those of eurypterids and arachnids, and are considered homologous. [2] The telson (the posteriormost division of the body) was small and semicircular, measuring 0.09 cm (0.036 in) in length. [1]
The appendages (limbs) were almost parallel and of a similar length, being uniform and pediform (foot-like). There was a small progressive increase in length until the marginally shorter fifth pair. The podomeres (segments of the appendages) were differentiated by a straight line and narrowed towards the distal end. The second and third pair of appendages had five distal podomeres (podomeres that were not underneath the prosoma) while the fourth and fifth, four. The sixth and last pair of appendages of Dvulikiaspis are among the best known in Chasmataspidida. They were modified into a "paddle" ubicated in front of the midsection of the prosoma. Although only five podomeres are known, it is possible that the sixth appendage of Dvulikiaspis had eight. The chelicerae (first pair of appendages) are not known. [1]
Dvulikiaspis was originally described in 1959 as a species of the eurypterid genus Stylonurus , S. menneri, based on one single nearly complete specimen, PIN 1271/2 (housed at the Paleontological Institute of the Russian Academy of Sciences, Moscow). It was found in deposits near the Imangda River at the southwest of the Taymyr Peninsula, near Norilsk, Krasnoyarsk Krai (Russia, then the Soviet Union). The specific name menneri honors the Russian paleontologist and geologist Vladimir Vasilyevich Menner, who discovered the fossil in 1956. He sent it to the Russian paleontologist Nestor Ivanovich Novojilov, who erected the species. [3] To date, his description is considered inaccurate, inexhaustive and based only on the best preserved material. [1] Novojilov reported the presence of paired tubercles in the center of the second and fourth to sixth tergites of S. menneri, and he compared this new species with the Scottish eurypterid Lamontopterus knoxae (then also part of Stylonurus). [3]
Based on these tubercles, Novojilov would later assign S. menneri to the eurypterid genus Tylopterella , with whom it shared this characteristic. The paired tubercles, however, were doubtfully present in the holotype. [2] Two new specimens, PIN 5116/1 (a nearly complete opisthosoma) and PIN 5116/3 (a poorly preserved dorsal fossil), were found in 1974 on an expedition by Yu. N. Mokrousov, but they were not described. They were found in the Upper Zub Formation-Lower Kureika Formation, in the Krasnoyarsk Krai. [1] In 2011, the British geologist and paleobiologist James C. Lamsdell noted that by the presence of a paddle-like sixth appendage, three-segmented fused buckler and nine-segmented postabdomen, T. menneri could in fact represent a chasmataspidid, questioning the position of T. menneri in Tylopterella and Eurypterida. [2]
In 2014, the paleontologists David J. Marshall, Lamsdell, Evgeniy S. Shpinev and Simon J. Braddy recognized T. menneri as a new separate genus of chasmataspidid due to its body size and position of the prosomal appendages, specially the sixth appendage in the anterior half of the prosoma. They also described the two specimens collected in 1974, determining that all known material of this genus was subject to taphonomic distortion (that is, defects product of the fossilization of the organism). They named it Dvulikiaspis, which is composed of the Russian word двуликий (dvulikij, "two-faced", referring to the long-lasting misidentification of D. menneri) and the Ancient Greek suffix ἀσπίς ( aspis , "shield"), being translated as "two-faced shield". They also noted that features such as the almost straight segment boundaries were shared with another chasmataspidid, Loganamaraspis , although the latter probably did not have a paddle-like sixth appendage. [1]
Dvulikiaspis is classified as part of the family Diploaspididae, one of the two families in the order Chasmataspidida. It includes one only species, D. menneri, from the Early Devonian of Siberia, Russia. [4]
D. menneri was originally recognized as a species of Stylonurus in 1959, [3] being moved to Tylopterella in 1962 due to the apparent presence of paired tubercles in the tergites. Both conclusions, done by Novojilov, have been criticized. Lamsdell already commented the possibility that D. menneri could represent a chasmataspidid in 2011. [2] This was demonstrated in 2014, when it was redescribed as a new genus of chasmataspidid and removed from Eurypterida, noting its distinction with other diploaspidids and suggesting a relationship with Loganamaraspis. [1] In 2019, during the description of the new Ordovician chasmataspidid Hoplitaspis hiawathai , it was suggested that Dvulikiaspis, Hoplitaspis and Loganamaraspis could represent a new family separated from Diploaspididae. These genera share the proportionality of the body (the postabdomen comprises the majority of the body length), a poorly differentiated buckler and a paddle projecting in front of the midpoint of the prosoma, although this is not certain in Loganamaraspis. However, the creation of a new family was not considered correct at the moment and all three genera currently remain in Diploaspididae, which is considered pending a more extensive revision. [5]
The cladogram below is based on a larger study (simplified to only show chasmataspidids) from 2015 carried out by the paleontologists Paul A. Selden, Lamsdell and Liu Qi, expanded to include the species Diploaspis praecursor , described in 2017. It showcases the phylogenetic relationships within Chasmataspidida. [6]
Chasmataspidida |
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Fossils of Dvulikiaspis have been discovered in Early Devonian (Lochkovian) deposits of Taymyria in Siberia, Russia. [1] D. menneri has been found alongside specimens of the chasmataspidids Heteroaspis stoermeri and Skrytyaspis andersoni , as well as the possible prosomapod Borchgrevinkium taimyrensis and indeterminate species of eurypterids like Acutiramus . The lithology (physical characteristics of the rocks) of the place near the Imangda River has been described as gypsiferous (with gypsum), dolomitic (with dolomite) and with dark gray marl. [3] [7] The fossils were collected 60 metres below the Early Devonian-Middle Devonian boundary. The lithology of the Upper Zub Formation-Lower Kureika Formation was not much different, also containing dolomitic marl with gypsum. The chasmataspidids remains were collected at between 307 m and 308 m. Here, D. menneri has been found with fossils of S. andersoni and the eurypterids Pterygotus and Parahughmilleria hefteri . [1]
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 million years ago. The group is likely to have appeared first either during the Early Ordovician or Late Cambrian period. With approximately 250 species, the Eurypterida is the most diverse Paleozoic chelicerate order. Following their appearance during the Ordovician, eurypterids became major components of marine faunas during the Silurian, from which the majority of eurypterid species have been described. The Silurian genus Eurypterus accounts for more than 90% of all known eurypterid specimens. Though the group continued to diversify during the subsequent Devonian period, the eurypterids were heavily affected by the Late Devonian extinction event. They declined in numbers and diversity until becoming extinct during the Permian–Triassic extinction event 251.9 million years ago.
Eurypterus is an extinct genus of eurypterid, a group of organisms commonly called "sea scorpions". The genus lived during the Silurian period, from around 432 to 418 million years ago. Eurypterus is by far the most well-studied and well-known eurypterid. Eurypterus fossil specimens probably represent more than 95% of all known eurypterid specimens.
Hibbertopterus is a genus of eurypterid, a group of extinct aquatic arthropods. Fossils of Hibbertopterus have been discovered in deposits ranging from the Devonian period in Belgium, Scotland and the United States to the Carboniferous period in Scotland, Ireland, the Czech Republic and South Africa. The type species, H. scouleri, was first named as a species of the significantly different Eurypterus by Samuel Hibbert in 1836. The generic name Hibbertopterus, coined more than a century later, combines his name and the Greek word πτερόν (pteron) meaning "wing".
Carcinosoma is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Carcinosoma are restricted to deposits of late Silurian age. Classified as part of the family Carcinosomatidae, which the genus lends its name to, Carcinosoma contains seven species from North America and Great Britain.
Chasmataspidids, sometime referred to as chasmataspids, are a group of extinct chelicerate arthropods that form the order Chasmataspidida. Chasmataspidids are probably related to horseshoe crabs (Xiphosura) and/or sea scorpions (Eurypterida), with more recent studies suggest that they form a clade (Dekatriata) with Eurypterida and Arachnida. Chasmataspidids are known sporadically in the fossil record through to the mid-Devonian, with possible evidence suggesting that they were also present during the late Cambrian. Chasmataspidids are most easily recognised by having an opisthosoma divided into a wide forepart (preabdomen) and a narrow hind part (postabdomen) each comprising 4 and 9 segments respectively. There is some debate about whether they form a natural group.
Alkenopterus is a genus of prehistoric eurypterid classified as part of the family Onychopterellidae. The genus contains two species, A. brevitelson and A. burglahrensis, both from the Devonian of Germany.
Nanahughmilleria is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Nanahughmilleria have been discovered in deposits of Devonian and Silurian age in the United States, Norway, Russia, England and Scotland, and have been referred to several different species.
Onychopterella is a genus of predatory eurypterid, an extinct group of aquatic arthropods. Fossils of Onychopterella have been discovered in deposits from the Late Ordovician to the Late Silurian. The genus contains three species: O. kokomoensis, the type species, from the Early Pridoli epoch of Indiana; O. pumilus, from the Early Llandovery epoch of Illinois, both from the United States; and O. augusti, from the Late Hirnantian to Early Rhuddanian stages of South Africa.
Willwerathia is a genus of Devonian arthropod. It is sometimes classified as synziphosurine, a paraphyletic group of horseshoe crab-like fossil chelicerate arthropods, while some studies compare its morphology to an artiopod. Willwerathia known only by one species, Willwerathia laticeps, discovered in deposits of the Devonian period from the Klerf Formation, in the Rhenish Slate Mountains of Germany.
Tylopterella is a genus of eurypterid, a group of extinct aquatic arthropods. Only one fossil of the single and type species, T. boylei, has been discovered in deposits of the Late Silurian period in Elora, Canada. The name of the genus is composed by the Ancient Greek words τύλη, meaning "knot", and πτερόν, meaning "wing". The species name boylei honors David Boyle, who discovered the specimen of Tylopterella.
Parahughmilleria is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of Parahughmilleria have been discovered in deposits of the Devonian and Silurian age in the United States, Canada, Russia, Germany, Luxembourg and Great Britain, and have been referred to several different species. The first fossils of Parahughmilleria, discovered in the Shawangunk Mountains in 1907, were initially assigned to Eurypterus. It would not be until 54 years later when Parahughmilleria would be described.
Unionopterus is a genus of eurypterid, an extinct group of aquatic arthropods commonly known as "sea scorpions". Fossils have been registered from the Early Carboniferous period. The genus contains only one species, U. anastasiae, recovered from deposits of Tournaisian to Viséan stages in Kazakhstan. Known from one single specimen which was described in a publication of Russian language with poor illustrations, Unionopterus' affinities are extremely poorly known.
Legrandella is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Legrandella was regarded as part of the clade Prosomapoda. Fossils of the single and type species, L. lombardii, have been discovered in deposits of the Devonian period in Cochabamba, Bolivia.
Weinbergina is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Fossils of the single and type species, W. opitzi, have been discovered in deposits of the Devonian period in the Hunsrück Slate, Germany.
Synziphosurina is a paraphyletic group of chelicerate arthropods previously thought to be basal horseshoe crabs (Xiphosura). It was later identified as a grade composed of various basal euchelicerates, eventually excluded form the monophyletic Xiphosura sensu stricto and only regarded as horseshoe crabs under a broader sense. Synziphosurines survived at least since early Ordovician to early Carboniferous in ages, with most species are known from the in-between Silurian strata.
Wiedopterus is a genus of eurypterid, an extinct group of aquatic arthropods. The type and only species of Wiedopterus, W. noctua, is known from deposits of Early Devonian age in Germany. The generic name derives from the Wied river, which runs near the site of the initial discovery, and the species name, noctua, derives from Latin noctua (owl) which refers to the superficial resemblance of the carapace to an owl.
Borchgrevinkium is an extinct genus of chelicerate arthropod. A fossil of the single and type species, B. taimyrensis, has been discovered in deposits of the Early Devonian period in the Krasnoyarsk Krai, Siberia, Russia. The name of the genus honors Carsten Borchgrevink, an Anglo-Norwegian explorer who participated in many expeditions to Antarctica. Borchgrevinkium represents a poorly known genus whose affinities are uncertain.
Camanchia is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Camanchia was regarded as part of the clade Prosomapoda. Fossils of the single and type species, C. grovensis, have been discovered in deposits of the Silurian period in Iowa, in the United States. Alongside Venustulus, Camanchia is one of the only Silurian synziphosurine with fossil showing evidence of appendages.
Anderella is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Anderella was regarded as part of the clade Prosomapoda. Fossils of the single and type species, A. parva, have been discovered in deposits of the Carboniferous period in Montana, in the United States. Anderella is the first and so far the only Carboniferous synziphosurine being described, making it the youngest member of synziphosurines. Anderella is also one of the few synziphosurine genera with fossil showing evidence of appendages, but the details are obscure due to their poor preservation.
Forfarella is a genus of chasmataspidid, a group of extinct aquatic arthropods. Fossils have been discovered in deposits of the Early Devonian period. The single and type species, F. mitchelli, is known from one only specimen found in Scotland, in the United Kingdom. Known as BMNH In 60023, it is poorly preserved and its type locality is uncertain, although it might be the Kelly Den stream section near the village of Arbirlot.